Overview of the Genus Phyllactinia (Ascomycota, Erysiphales) in Azerbaijan
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Plant & Fungal Research (2018) 1(1): 9-17 © The Institute of Botany, ANAS, Baku, Azerbaijan http://dx.doi.org/10.29228/plantfungalres.2 December 2018 Overview of the genus Phyllactinia (Ascomycota, Erysiphales) in Azerbaijan Dilzara N. Aghayeva1 Key Words: distribution, ectoparasitism, endoparasit- Lamiya V. Abasova ism, host plant, plant pathogen, powdery mildews Institute of Botany, Azerbaijan National Academy of Sciences, Badamdar 40, Baku, AZ1004, Azerbaijan INTRODUCTION Susumu Takamatsu Graduate School of Bioresources, Mie University, Powdery mildews are one of the frequently encoun- 1577 Kurima-Machiya, Tsu 514-8507, Japan tered plant pathogens and most of them are epiphytic (14 genera from 18), in which they tend to produce hy- Abstract: Intergeneric diversity of powdery mildews phae and reproductive structures on surface of leaves, within the genus Phyllactinia in Azerbaijan was inves- young shoots and inflorescence [Braun, Cook, 2012; tigated using morphological approaches based on re-ex- Glawe, 2008]. These fungi absorb nutrients from plant amination of herbarium specimens kept in Mycological tissue via haustoria, which develops in epidermal cells Herbarium of the Institute of Botany (BAK), Azerbaijan of plants [Braun, Cook, 2012]. Among all powdery mil- National Academy of Sciences and collections of re- dews only four genera demonstrate endoparasitism, of cent years. To contribute detail taxonomic analysis data them Phyllactinia Lév., have partly endoparasitic na- given in literatures was revised. Modern taxonomic and ture. Fungi belonging to these genera penetrate into the nomenclature changes were considered. The host range plant cell via stomata and develop haustoria in paren- and geographical distribution of species residing to the chyma cells. Endoparasitic genera of powdery mildews genus within the country were clarified. Consequently, (Phyllactinia, Pleochaeta Sacc. & Speg., Queirozia 17 powdery mildew taxa were recorded, of which Ph. Viégas & Cardoso and Leveillula G. Arnaud) form a alnicola, Ph. ampelopsidis, Ph. babayanii, Ph. carpini, distinct monophyletic group within Erysiphales which Ph. corni, Ph. fraxini, Ph. guttata, Ph. mali, Ph. ma- suggests that endoparasitism derived from ectoparasit- rissalii, Ph. moricola, Ph. nivea, Ph. orbicularis, Ph. ism in powdery mildews only once in their evolution paliuri, Ph. phaseolina, Ph. populi, Ph. pyri-serotinae [Takamatsu, 2013; Takamatsu et al. 2016]. Only genus and Ph. roboris on 32 plant species from 11 families Leveillula exhibits true endoparasitism and elongates were listed. These families belong to the groups Aster- internal hyphae between host cells in mesophyll layer ids (Cornaceae, Oleaceae) and Rosids (Fabaceae, Betu- [Takamatsu et al. 2008, 2016]. laceae, Fagaceae, Salicaceae, Moraceae, Rhamnaceae, Phyllactinia is distinguishable from other powdery Rosaceae, Ulmaceae, Sapindaceae, Vitaceae), of which mildews based on its unique characteristics of sexual Rosids exceeds in number of host species. The family morphs, viz. large ascomata (chasmothecia) with two Rosaceae possess the large amount of host species, but different types of appendages. One of them is acicular specific affinity of genus Phyllactinia is considered to appendages with bulbous swellings at the base, which be to the family Betulaceae. In addition, Phyllactinia arise equatorially and play role in orientation of asco- powdery mildews seemed to spread mainly in moun- mata through the air during dispersal. Second one, peni- tainous areas of Azerbaijan. The highest number of taxa cillate cells composed of foots (outgrows of the asco- was recorded in the districts belonging to the Southern- matal walls), filaments [Shin, Lee, 2002] and grouped East Great Caucasus region of the country, followed by together on the dorsal surface of chasmothecia [Glawe, regions Middle Araz and Lesser Caucasus. The low- 2008]. Penicillate cells perform a role as anchor for at- est number of Phyllactinia taxa was recorded from taching of chasmothecia to the new plant surface by Kur lowland and Lankaran regions. During the present gelatinizing of filaments under humid condition [Taka- study, it was revealed that herbarium samples of 14 taxa matsu et al., 2008]. There is a speculation that unique of genus Phyllactinia are available in the BAK, infor- characteristics of appendages are an evolutionary adap- mation of which is provided in this article. tation to the tree parasitism of this genus [Takamatsu et al., 2008]. Powdery mildews from this genus form Accepted for publication: 5 December 2018 whitish to grayish colony under the leaf surface. The 1E-mail: [email protected] genus is characterized by having 2–3-spored asci, main- 9 Plant & Fungal Research ly monomorphic, sometimes dimorphic conidia with clavate, dumble-like, subcylindric, broadly ellipsoid- ovoid, lanceolate shape, almost indistinct and nipple or rod-shaped appressoria [Braun, Cook, 2012] (Fig. 1). According to the monograph of the Erysiphales, 102 species were counted within the genus Phyllactinia [Braun, Cook, 2012] and they are strictly woody para- sitic fungi, which are known on about 700 plant species from 69 families [Amano, 1986]. Totally, 108 plant spe- cies from family Rosaceae were dedected as host for genus Phyllactinia and the order is followed by families Betulaceae (91 species), Fagaceae (56 species), Oleace- ae (50 species), etc. [Takamatsu et al., 2008]. The aims of this study were to reveal diversity of powdery mildews from the genus Phyllactinia, to clar- ify the ratio of Phyllactinia taxa within the Erysiphales in Azerbaijan, to update species composition by taking Figure 1. Phyllactinia moricola; A. acicular appendage into the consideration modern taxonomic and nomen- with bulbous swelling; B. penicillate cell, b1. foot, b2. clature changes, and to investigate their host range and filaments; C. conidia; D. conidiophores; E. appressoria; geographical distribution within country. Bar = 100 μm. MATERIAL AND METHODS Herbarium samples kept at the Mycological Herbari- RESULTS AND DISCUSSION um of the Institute of Botany, ANAS (BAK) and new Taxonomy. Diversity of powdery mildews in Azerbaijan samples collected from 2016 to 2018 were investigated is currently represented with 134 fungal taxa from ten using classic morphological approaches. To contribute genera on 420 plant species from 51 families [Abasova, detail taxonomic investigations, literature data were Ağayeva, 2018]. The genus Erysiphe R. Hedw. ex DC. summarized. dominates by possessing the largest number of fungal Morphological examinations were done under the taxa, followed by the genus Podosphaera Kunze, Go- optical microscope (Vert. A1, Carl Zeiss, Axio Imager, lovinomyces (U.Braun) Heluta, Phyllactinia, and Lev- Göttingen, Germany). To examine asexual structures a eillula. Genera Arthrocladiella Vassilkov, Blumeria little piece of infected leaf was rehydrated using lactic Golovin ex Speer, Neoërysiphe U.Braun and Sawadaea acid method [Shin, La, 1993]. For observation of sexual Miyabe are quite small and consist of one or two species morphs, chasmothecia were mounted in 3% NaOH so- for each genus [Abasova, Ağayeva, 2018]. Intergeneric lution [Meeboon, Takamatsu, 2015]. Thirty measure- diversity of the genus Phyllactinia, their host range, ments for each structure were taken. Measurements geographical distribution within the country, and tradi- of asexual structures were done based on the length of tional names put on the specimens and BAK numbers of conidiophores, size and shape of conidia and appres- were given below. soria. For sexual structures diameter of chasmothecia, Ph. alnicola U. Braun, Braun & Cook 226, 2012; and number, size and shape of asci and ascospores were = Ph. suffulta f. alni Hammarl.; Host: Alnus glutinosa considered. Bulmer’s factor (l × 1.5; w × 1.2) was used (L.) Gaertn. (Betulaceae); BAK 5612, 5614, 5615; Dis- to reconstruct the original size of conidia for herbarium tribution: Qusar, Zaqatala (Fig.2.1). samples [Braun, Cook, 2012]. Drawings were made by Ph. ampelopsidis Y.N. Yu & Y.Q. Lai, Acta Micro- freehand using scale bar. biol. Sin. 19 (1): 14, 1979; Host: Vitis vinifera L. (Vi- Identification of fungal species was carried out by taceae); BAK 5778; Distribution: Khojavend. Previ- using the latest literature [Braun, Cook, 2012]. Nomen- ously a powdery mildew fungus on Vitis vinifera from clature changes were done according to the Mycobank Azerbaijan was described as Phyllactinia suffulta f. vi- and Index Fungorum. Systematic assignment of plants tis Jacz. [Huseynova, 1961a]. Herbarium specimen was was performed based on APG IV system [APG IV, examined morphologically in this study, but not any 2016]. other structure characterizing genus Phyllactinia was 10 Aghayeva et al.: The genus Phyllactinia in Azerbaijan observed. Description of Ph. suffulta f. vitis provided identified asPh. fraxini. by B. Huseynova is quite similar to those of Ph. am- Ph. guttata (Wallr.: Fr.) Lév., Bot., 3 Sér., 15: 144, pelopsidis in 180–200 μm diameter of chasmothecia [in 1851; ≡ Ph. corylea (Pers.) P.Karst.; = Ph. suffulta f. monograph 150–280 μm], 6–8 number [5–14] and up to coryli-avellanae Jacz.; Host: Corylus avellana L., C. 250 μm length [250–370 μm] of acicular appendages, colurna L. (Betulaceae); BAK 5597–5601, 5637–5645, 54–76 × 22–30 μm [50–100 × 20–45 μm] and ellipsoid- 5648, 5650–5658; Distribution: Qusar, Khachmaz, ovoid, 2(–3)-spored of asci. However, Ph. suffulta f. Shabran, Shaki, Oghuz, Qakh, Zaqatala, Lankaran, vitis was given the affinity